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Materials
and methods
Results
Discussion
References
The dissemination of Striga hermonthica (buda) seed by animals may occur through three processes. First, fresh plants bearing mature seed may be fed to domestic animals as a component of forage feed; second, it may be grazed in the field after the sorghum crop has been harvested; and third, it may become attached to animals' coats while the animals are resting in or traversing infested aura fields. Hansen (1911) reported that a cow grazing on a weedy field consumed in a day 89,000 seeds of Plantago spp. and 564,000 seeds of Matricaria chamomilla, of which 85,000 and 198,000 respectively were voided at 58 per cent and 27 per cent germination capacity.
Since the host sorghum {aura) and its specific parasitic weed buda co-occur in field situations and are utilized by domestic animals as a mixture for forage, either fresh or dry, and because buda has become the predominant weed affecting aura, the present study was conducted to furnish information on the biotic relations of domestic animals and buda in regard to its dissemination and utilization. The study aims to answer the following three questions: - What is the potential role of animal dung in buda seed dissemination? - What is the effect of animal digestive juices on buda germination capacity and relative effects on buda dissemination? - What is the potential of fresh buda plants as a feed component?
In an attempt to find answers to these questions three experiments were carried out. The first two experiments were conducted to investigate the role of ruminants in the dissemination of buda. The third experiment evaluated buda's nutritive-value. Experiment 1 involved evaluation of budainfested sorghum fed to animals, and experiment 2, the response of buda seed to incubation in rumen liquor in vivo and in vitro.
Experiment 1
Three animals - a goat, a sheep, and a camel - were fed for four weeks, commencing 1 September 1979, on forage sorghum (Sorghum bicolor var. Abu Sabeen) which was heavily invested with buda. Faeces of each animal were collected daily and spread on hard cloth material under the sun to dry. Two weeks after feeding was discontinued, the sun dried faeces were collected and ground into powder using a large local wooden mortar. Subsamples of faeces 100 9 each were used as inoculum material. Clay pots (12 cm diameter) were filled with 2 kg of river silt to which the inoculum material was added as required. There were four treatments, using soil inoculated with (a) infested goat faeces, (b) infested sheep faeces, (c) infested camel faeces, and (d) no faeces (control). Each pot was sown with eight seeds of sorghum, which were later thinned to four seedlings per pot. A randomized complete block design with four replications was used. Statistical analysis was performed on transformed data (square roots).
Experiment 2
Seeds of three buda populations - from Shambat, Medani, and Abu Na'ama - were collected in November 1979, and in August 1980 were subjected to incubation periods in rumen liquor in vivo and in vitro to test the effect on their germination capacity.
In the in vivo part of the experiment, 0.01 9 samples of each seed population were wrapped in 2 x 2 cm squares of Nitex material, which were then tied firmly into bags using synthetic plastic string with a lead of about 30 cm. The bags were suspended in the rumen of a fistulated sheep for different periods of time. There were five treatments: incubation for one day, incubation for two days, incubation for four days, incubation for eight days, and no incubation (control).
At the end of each treatment, the seeds were removed from the bags and washed thoroughly with distilled water and then were sterilized by soaking in a 1 per cent chlorine solution (as sodium hypochlorite) for five minutes. They were then washed thorouqhiv again and left to dry overnight on Whattman filter paper.
Each sample was divided into six portions placed on separate discs, with approximately 30 seeds per disc. The seeds were then set up for preconditioning at 23 C on moist giass-fibre filter paper as described by Parker et al. (1977).
The discs of preconditioned buda seeds were dabbed on dry filter paper to remove excess moisture and placed in groups of six in a petri dish which contained glass-fibre filter paper moistened with 5 ppm of GR 45 (synthetic germination stimulant). The dishes were then closed, wrapped in polyethylene bags, and incubated at 33 C for 24 hours. After this incubation period the germinated seeds were counted.
The in vitro part of the experiment was carried out under laboratory conditions using Tilley and Terry's (1963) technique for in vitro digestion of forage crops. Statistical analyses were performed on transformed data (arcsin).
Experiment 3
An air-dried sample of buda was taken from a random sample collected at different times of the growing season and from different geographical locations within Sudan. The plant was chopped into small pieces and ground in a Christy Norris mill using a 0.8 mm sieve. The dry matter of the weed was then determined and the proximate analysis was carried out according to the official methods of analysis of the Association of Official Agricultural Chemists (AOAC 1956).
Buda was also fed to six lambs (Sudan desert-sheep) for a period of 17 days in order to determine its digestibility and nitrogen retention. During the experimental period the animals were housed in metabolic crates especially designed for total urine collection. Faeces collection was carried out by canvas bags fitted to the animals by harnesses.
The material under investigation was offered ad libitum, and free access to water was maintained. The experiment started with a 10day preliminary period in which no faeces or urine collections were made. This period was to ensure the adaptation of the animals to the harnesses, bags, and crates and of the rumen microbes to the feed being consumed by the animals (Lloyd et al. 1956). The feed was offered once a day, at 8.00 a.m. Any residues from the previous day were collected at 7.30 a.m., dried, and weighed, and the amount was deducted from the drymatter weight of food provided. Daily subsamples of the food being offered and of the residues of each animal were treated in a manner similar to that described by El-Hag (1976). . and urine from each animal were collected separately each day before providing the day's ration and received a treatment similar to that described by Abou Akkada and El-Shazly (1965).
The dried samples of the residues and faeces were ground in the mill mentioned above, and the samples were redried before weighing for analysis. Total nitrogen in the urine was estimated by the micro-Kjeldahl method used by El-Shazly (1958).
On the last day of the experimental period, samples of rumen liquor were obtained by means of a stomach tube. The first sample was collected at 8.00 a.m., just before introducing the feed, and the second about three hours after feeding. The rumen liquor was strained through two layers of gauze and kept for immediate analysis. Total VFA and ruminal ammonia nitrogen were determined by the method explained by Abou Akkada and El-Shazly (1958, 1965).
TABLE 10. Mean number of buda (Striga hermonthica) plants per pot
Treatment | Number |
Control | 0 |
Sheep | 1.653 |
Goat | 2.978 |
Camel | 1.045 |
Experiment 1
The effects of infested animal dung on the dissemination of buda are shown in table 10. Highly significant differences (P = 0.01) of buda incidence were observed between treatments. Infested goat dung induced the greatest buda infestation, and infested camel dung the lowest. No significant differences of buda incidence were found between camel and control treatments. However, highly significant differences (P = 0.01) of infestation existed between goat and control or sheep treatments.
Experiment 2
Table 11 shows the effects of incubation periods in rumen liquor under in vivo conditions. Taking all the buda seed populations together, the greatest germination response was observed in the control treatment and the lowest in treatments where the seed were incubated in the sheep's rumen for a period of four or eight days. Highly significant differences (P = 0.01) existed between the treatments. Again taking all the strains together, a significant reduction in germination capacity was manifested when the seed were incubated in rumen liquor for one or two days, with complete loss of seed viability over a period of between approximate three and eight days. No significant differences in germination response was manifested between seeds incubated for one day and those incubated for two days.
Highly significant differences (P = 0.01) in strain x treatment interaction were observed (table 11). Seeds of the population from Medani scored the highest germination response, and those from Shambat the lowest. The Medani strain, in contrast to those from Abu Na'am and Shambat, appeared to respond favourably to the effects of rumen liquor, resulting in an increased germination response after both one-day and two-day incubation periods.
Data from the in vitro part of the experiment generally agree with the data of the in vivo part with two exceptions (table 12). First, the period required for complete loss of seed viability of buda seed populations was reduced to approximately four days. Second, with regard to the strain x treatment interaction, although the strains responded differently to the incubation period in rumen liquor, most of them responded unfavourably with consequent reductions of germination capacity.
TABLE 11. Germination response of buda as affected by different incubation period in rumen liquor in vivo
Buda population | Treatment | Population mean% | ||||
C | 1 | 2 | 4 | 8 | ||
Shambat | 122.2 | 81.0 | 85.9 | 0 | 0 | 9.6 |
Medani | 277.5 | 308.3 | 369.9 | 0 | 0 | 31.8 |
Abu Na'ama | 323.8 | 227.6 | 119.3 | 0 | 0 | 22.3 |
Treatment mean % | 30.1 | 34.2 | 31.9 | 0 | 0 |
C = control (not incubated); 1, 2, 4, 8 = days of incubation
LSD at P = 0.05 for:
buda populations | = 2.3 |
incubation treatments | = 2.8 |
buda x treatment interaction | = 4.8 |
Experiment 3
The proximate composition of the sample of buda analysed is shown in table 13. The buda was found to be rich in crude protein and relatively low in crude fibre in comparison to roughage feeds. The ash content seemed to be high, ranging between 20 and 23 per cent.
All the values given in tables 14-16 are means of the results from the six animals used. Table 14 shows the apparent digestibility of the dry matter and its constituent nutrients: the dry matter had a digestibility of 47.0 per cent, and the crude protein and crude fibre were 55.1 and 51.3 per cent digestible respectively.
The nutritive value expressed in terms of total digestible nutrients (TON), starch equivalent (SE), digestible crude protein (DCP), and nitrogen balance is shown in table 15. The buda had a TON value of 41.4 and a starch equivalent of 29.0; its digestible crude protein amounted to 6.94 and the nitrogen balance to 4.4 9 N.
Table 16 shows that the mean change in the concentration of VFA in the rumen of the sheep over the period from just before they were fed buda until three hours afterwards was 3.9 m-equivalent/100 ml rumen liquor. The change in the concentration of rumen ammonia over the same period of three hours was 6.4 mg N/100 ml rumen Iiquor.
The findings of the present study suggest that domestic animals may act as potentially effective distributors of seeds of the parasitic weed buda (Striga hermonthica). There is evidence that these seeds may be transported through animal dung.
The general contention is that storm water and wind are the chief means of spreading the seeds, but Farquahar (1937) stated that cattle are the chief means of spreading. He noticed that witchweed (Striga spp.) spread more rapidly in Rhodesia than in Natal; the reason attributed was that in Natal the pest was killed off early by heavy frosts. In an experiment, Farquahar also noticed that in every field that had been shoocked, there was an increase in the pest the following season. Lands were cleaned up faster when cattle were not allowed in cultivated lands. In another experiment, he compared the dung from cattle fed solely on grasses and from cattle fed a small quantity of the weed. The dung was applied to soil containing maize seeds. Witchweed appeared profusely in boxes which received dung from cattle that had been fed on witchweed plants and none appeared in the other.
The present findings provide additional evidence suggesting that the effects of digestive juices of ruminant animals may affect buda seeds either adversely or favourably and that this in turn depends on the seed strain and also on the incubation period of the seed in rumen liquor. The adverse effects may range from reduction of germination capacity to complete loss of seed viability.
Over all, the adverse effects of digestive juices of ruminants in both in viva and in vitro experiments is indicated by the significant reduction of germination capacity of buda seeds, with approximately 14 and 20 per cent below the control at one or two days' incubation period in vivo and approximately 8, 37, or 76 per cent below the control at one, two, or four days' incubation period in vitro. However, not all striga strains responded adversely to an increased incubation period in rumen liquor, the exception being the Medani strain, which had an increase of germination response of approximately 11 per cent and 33 per cent over the control at one or two days' incubation period in rumen liquor in vivo.
TABLE 12. Germination response of buda as affected by different incubation periods in rumen liquor in vitro
Buda population | Treatment | Population mean % | ||||
C | 1 | 2 | 4 | 8 | ||
Shambat | 122.2 | 64.2 | 42.6 | 0 | 0 | 7.6 |
Medani | 277.5 | 254.2 | 201.5 | 77.5 | 0 | 27.0 |
Abu Na'ama | 323.8 | 347.3 | 211.3 | 89.8 | 0 | 32.4 |
Treatment mean % | 24.1 | 22.1 | 15.1 | 5.5 | 0 |
C = control (not incubated; 1, 2, 4, 8 = days of incubation
LSD at P = 0.05 for:
buds populations | = 1.4 |
incubation treatments | = 3.2 |
buda x treatment interaction | = 5.5 |
TABLE 13. Chemical composition of S. hermonthica on dry-matter basis (percentages)
Dry matter | 93.6 |
Crude protein | 12.6 |
Crude fibre | 20.3 |
Ether extract | 1.7 |
Nitrogen-free extract | 44.1 |
Ash | 21.3 |
TABLE 14 Mean digestibility coefficients for the components of S. hermonthica
Dry matter | 47.0 |
Organic matter | 51.5 |
Crude protein | 51.1 |
Crude fibre | 51.3 |
Ether extract | 41.7 |
Nitrogen-free extract | 51.0 |
TABLE 15. Nutritive value of S. hermonthica
Total digestible nutrients | 41.4 |
Starch equivalent | 29.0 |
Digestible crude protein | 6.9 |
Nitrogen retained (g) | 4.4 |
TABLE 16. Change in the concentration of VFA (mequivalent/100 ml rumen liquor) and ruminal ammonia (mg N/100 ml rumen liquor)
VFA | Ammonia | |
Concentration at 8 a.m. | 4.47 | 8.21 |
Concentration at 11 a.m. | 8.37 | 14.61 |
Change | +3.9 | +6.4 |
It is interesting to note that incubation of buda seeds in rumen liquor for more than two days in viva may lead to complete loss of seed viability. Thus the time taken for the seeds to pass through the digestive tract of ruminants appears to be a critical factor in the survival and distribution of the seeds. It is known that the rate of excretion of animal faeces depends to a large extent on the fluid content of the digestible matter. Consequently it may be assumed that the percentage of moisture in the feed stuff or the quantity of drinking water available or the drinking interval may influence the process of digestion and the time taken for digestible material to pass through the digestive tract of animals. Therefore, it may be assumed that in field situations under irrigation domestic animals fed on buda. infested forage may excrete dung within 8 to 24 hours. This may be conducive to relatively less damage to germination capacity than in field situations under rain-fed conditions where the intervals between drinking are prolonged because of the distances between and limited availability of water sites. However, there are villages in rain-fed areas whose domestic animals go out early in the morning to graze and return later in the evening to drink. These animals of course do not experience the long intervals between drinking and therefore constitute a major hazard leading to buda dispersal, especially in the vicinity of the villages.
The high percentage of crude protein in buda obtained in this study is of particular importance in a tropical country like Sudan, where most of the roughagesare deficient in this important nutrient. This relative abundance of crude protein coupled with the low crude fibre content brings the buda parasite nearer to the class of concentrates than roughages. One of the striking features of the composition of buda was its high ash content (21.3 per cent), which was appreciably higher than that found in normal feedstuffs. The richness of this plant material in ash could not be attributed to roil contamination as all the different samples analysed indicated more than 20 per cent ash content.
The proximate analysis that appears in table 13 provides information about the potential value of the material as a feed, but the actual value of a feed to the animal can be determined only after feeding it to the animal and making allowance for the inevitable losses that occur during the utilization of the feed. The most important of these losses is that represented by the part that is not absorbed and is excreted in the faeces. The apparent digestibility measurement is therefore the most important factor in determining the value of the feed.
The apparent digestibility of the buda dry matter was found to be 47 per cent, which is significantly lower than the in vitro digestibility value of 58 per cent we reported in an earlier study (Bebawi et al. 1980). The discrepancy between the two digestibility values might be mainly attributed to the difference in the chemical composition of the two samples of buda evaluated. It is known that the crude fibre percentage increases with age and maturity and as the crude fibre increases, digestibility decreases. The material used in the in vivo study might have been more mature than that used in the in vitro study. It was noticed that the apparent digestibility of the buda dry matter was lower than that obtained for the most common foodstuff available in Sudan, bummera (desert grasses), (El-Hag 1976). However the TDN value, starch equivalent, and digestible crude protein were higher for the buda than for hummra (41.4 vs. 31.4, 29.0 vs. 21.1, and 6.1 vs. 1.6 for TDN, SE, and pop for buda and hummra respectively).
The production of VFA, the ultimate metabolic products of feed degradation in the rumen, is a good indicator of the activity of the rumen micro-organisms. The 3.9 mequivalent/100 ml rumen liquor change in the concentration of VFA compared favourably to that obtained with most foodstuffs found in Sudan. This confirmed that the activity of the micro-organisms was not affected as a result of feeding buda.
The high concentration of rumen ammonia three hours after feeding indicated that a great proportion of buda nitrogen was not utilized for useful purposes by the animal as it is well established that there is a close negative correlation between ammonia concentration in the rumen and the usefulness of dietary protein to the animal (Annism et al. 1954; Chalmers and Synge 1954). This low utilization of protein might be attributed to a shortage of soluble carbohydrates in buda, as without this nutrient the ammonia produced in the rumen cannot be trapped by the microorganisms to synthesize microbial protein. An implication of this finding is that buda should be fed together with other foodstuffs rich in soluble carbohydrates in order to improve nitrogen utilization.
Measurements of nitrogen balance are considered more useful than ammonia concentration in assessing the value of protein to ruminant animals, but the amount of work usually involved is far greater. The average amount of nitrogen retained by the animals was 4.43 9, a figure that reflected an efficient utilization of the crude protein, especially if we know that the nitrogen retained by sheep fed on Medicago stiva was only 3.4 9 (El-Hag 1976). The results of the nitrogen balance measurement seemed not to be in line with the high ammonia concentration found in the rumen. However, there is always difficulty in measuring the N-balance since it is affected by the pre-experimental history of the animal and the duration of the experimental period.
Buda was readily accepted by the sheep and produced no adverse influence on rumen fermentation. It was evident that it contained no toxic material at the level taken by the animals as none of them showed any ill effects from its consumption.
It may be concluded from the findings of the present study that domestic animals may be one of the chief distributors of S. hermonthica (buda) seed in areas both of irrigated and of rain-fed agriculture in Sudan. Howver, it is expected that relatively more efficient distribution of the seed may be achieved by animals in areas of irrigated agriculture or by animals belonging to settled villagers in rain-fed areas than by those owned by pastoralists in rain-fed areas.
It could also be concluded that buda is a potentially useful feedstuff and could safely be considered as a mediumquality roughage. Since it grows in association with aura, which is usually grown in summer in Sudan, the weed will be available at a time of great feed scarcity, and this adds to its potential importance as a feed. The question of using buda as a feed is therefore worth further investigation. Two points should be noticed, however: (a) buda not only leads to the reduction of the aura crop yield but also stunts the stem of the aura, and it is not yet clear whether the gain in buda as a fodder would compensate for the reduction of stubble which is also used as feed for the animals; and (b) this method of using the buda plant can only really be effective if quick-maturing varieties of aura can prove themselves popular and thus allow farmers to resort to trap-cropping, so that the products can be used as fodder and at the same time help reduce the level of buda infestation.
In summary, the existence of Striga hermonthica (buda) in the arid and semi-arid region of Sudan has acted as a deterrent to the adoption of improved husbandry methods. Methods of control so far developed are too complicated for the illiterate farmer to comprehend and too costly for the poorer peasants. At the same time severe questions about the efficacy of many control measures makes many of the better-off farmers in the modern sector doubt the economic feasibility of such methods. It has been demonstrated that buda can be successfully used as a feed in association with other feeding stuffs, but the price to be paid seems likely to be an increase in buda infestation, especially in irrigated areas and rain-fed areas with a regular daily watering pattern. Nomadic animals may be less of a hazard than previously suggested because of the loss of buda seed viability with increased length of immersion in rumen liquor. Clearly more research and a coherent programme of agricultural extension among farmers on buda is required before this obstacle to the adoption of better husbandry in the semi-arid and arid lands of Sudan can be overcome.
H. B. J. Davies
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